Kenneth H. Pollock

Design and analysis methods for fish survival experiments based on release-recapture

Design and analysis methods for fish survival experiments based on release-recapture / , Design and analysis methods for fish survival experiments based on release-recapture / , مرکز فناوری اطلاعات و اطلاع رسانی کشاورزی

Survival analysis in telemetry studies: The staggered entry design

The estimation of survival distributions for radio-tagged animals is important to wildlife ecologists. Allowance must be made for animals being lost (or censored) due to radio failure, radio loss, or emigration of the animal from the study area. The Kaplan-Meier procedure (Kaplan and Meier 1958), widely used in medical studies subject to censoring, can be applied to this problem. We developed a simple modification of the Kaplan-Meier procedure that allows for new animals to be added after the study has begun. We present 2 examples using telemetry data collected from northern bobwhite quail (Colinus virginianus) to show the simplicity and utility of the Kaplan-Meier procedure and its modifications. The log rank test used to compare 2 survival distributions can also be modified to allow for additions during the study. Simple computer programs that can be run on a personal computer are available from the authors. J. WILDL. MANAGE. 53(1):7-15 Radio-tagged animals are used to study survival. Present techniques for analyzing data from these studies assume that each survival event (typically an animal surviving a day) is independent and has a constant probability over all animals and all periods (Trent and Rongstad 1974, Bart and Robson 1982, Heisey and Fuller 1985). We believe these assumptions are often unrealistic and restrictive. White (1983) generalized discrete approaches using the same framework as that of band return models (Brownie et al. 1985) and he developed a flexible computer program (SURVIV) for use with his approach. Heisey and Fuller (1985) generalized the Trent and Rongstad (1974) approach to allow mortality from different causes (e.g., predation, starvation) and developed a microcomputer program called MICROMORT. Typically an animals exact survival time (at least to within 1-2 days) is known unless that survival time is right censored (i.e., only known to be greater than some value). Pollock (1984) and Pollock et al. (1989) suggested a useful approach based on continuous survival models allowing right censoring that is widely used in medicine and engineering (Kalbfleisch and Prentice 1980, Cox and Oakes 1984) and provided examples of the Kaplan-Meier procedure. The Kaplan-Meier procedure does not require specification of a particular parametric continuous distribution; e.g., the exponential or Weibull. Related ecological papers using survival methods include Muenchow (1986), Pyke and Thompson (1986), Kurzejeski et al. (1987), and White et al. (1987). We present a simple description of the Kaplan-Meier procedure with an example using northern bobwhite quail survival data collected by PDC. We then generalize the Kaplan-Meier procedure to allow gradual (or staggered) entry of animals into the study. The calculations are illustrated with an example from the quail data. Finally, we present the log-rank test for comparison of survival distributions (modified for staggered entry of animals) with an example. We also present a discussion of model assumptions and directions for future research. We thank J. D. Nichols and W. L. Link for helpful comments on an earlier draft of this paper. We acknowledge G. C. White and D. M. Heisey for their helpful reviews that improved the final version. THE KAPLAN-MEIER OR PRODUCT LIMIT PROCEDURE The Kaplan-Meier or product limit estimator was developed by Kaplan and Meier (1958) and is d scussed by Cox and Oakes (1984:48) and Kalbfleisch and Prentice (1980:13). The survival function (S[t]) is the probability of an arbitrary animal in a population surviving t units of time from the beginning of the study. A nonparametric estimator of the survival function can be obtained by restricting ourselves to the discrete time points when deaths occur a1, a2, ..., ag. We define r, . . . , rg to be the numbers of an-

ESTIMATING SPECIES RICHNESS: THE IMPORTANCE OF HETEROGENEITY IN SPECIES DETECTABILITY

Estimating species richness (i.e., the actual number of species present in a given area) is a basic objective of many field studies carried out in community ecology and is also of crucial concern when dealing with the conservation and management of biodiversity. In most studies, the total number of species recorded in an area at a given time is taken as a measure of species richness. Here we use a capture–recapture approach to species richness estimation with North American Breeding Bird Survey (BBS) data in order to estimate species detectability and thus gain insight about its importance. In particular, competing models making different assumptions about species detectability are available. We carried out analyses on all survey routes of four states, Arizona, Maryland, North Dakota, and Wisconsin, in two years, 1970 and 1990. These states were chosen to provide contrasting habitats, bird species composition, and survey quality. We investigated the effect of state, year, and observer ability on the propo...

A REMOVAL MODEL FOR ESTIMATING DETECTION PROBABILITIES FROM POINT-COUNT SURVEYS

Abstract Use of point-count surveys is a popular method for collecting data on abundance and distribution of birds. However, analyses of such data often ignore potential differences in detection probability. We adapted a removal model to directly estimate detection probability during point-count surveys. The model assumes that singing frequency is a major factor influencing probability of detection when birds are surveyed using point counts. This may be appropriate for surveys in which most detections are by sound. The model requires counts to be divided into several time intervals. Point counts are often conducted for 10 min, where the number of birds recorded is divided into those first observed in the first 3 min, the subsequent 2 min, and the last 5 min. We developed a maximum-likelihood estimator for the detectability of birds recorded during counts divided into those intervals. This technique can easily be adapted to point counts divided into intervals of any length. We applied this method to unlimited-radius counts conducted in Great Smoky Mountains National Park. We used model selection criteria to identify whether detection probabilities varied among species, throughout the morning, throughout the season, and among different observers. We found differences in detection probability among species. Species that sing frequently such as Winter Wren (Troglodytes troglodytes) and Acadian Flycatcher (Empidonax virescens) had high detection probabilities (∼90%) and species that call infrequently such as Pileated Woodpecker (Dryocopus pileatus) had low detection probability (36%). We also found detection probabilities varied with the time of day for some species (e.g. thrushes) and between observers for other species. We used the same approach to estimate detection probability and density for a subset of the observations with limited-radius point counts.

ESTIMATING SITE OCCUPANCY AND SPECIES DETECTION PROBABILITY PARAMETERS FOR TERRESTRIAL SALAMANDERS

Recent, worldwide amphibian declines have highlighted a need for more extensive and rigorous monitoring programs to document species occurrence and detect population change. Abundance estimation methods, such as mark–recapture, are often expensive and impractical for large-scale or long-term amphibian monitoring. We apply a new method to estimate proportion of area occupied using detection/nondetection data from a terrestrial salamander system in Great Smoky Mountains National Park. Estimated species-specific detection probabilities were all <1 and varied among seven species and four sampling methods. Time (i.e., sampling occasion) and four large-scale habitat characteristics (previous disturbance history, vegetation type, elevation, and stream presence) were important covariates in estimates of both proportion of area occupied and detection probability. All sampling methods were consistent in their ability to identify important covariates for each salamander species. We believe proportion of area occupie...

A Review of Tagging Methods for Estimating Fish Population Size and Components of Mortality

Abstract Techniques to improve estimation of animal population size and mortality from tagging studies have received substantial attention from terrestrial biologists and statisticians during the last 20 years. However, these techniques have received little notice from fisheries biologists, despite the widespread applicability to fisheries research, the wide variety of tag types used in fisheries research (from traditional fin clips to telemetry tags), and the development of new computer software to assist with analyses. We present a brief review of population models based on recaptures, returns, or telemetry relocations of tagged fish that can be used to estimate population size, total mortality, and components of mortality (i.e., fishing and natural) that are frequently of interest to fisheries biologists. Recommended strategies include (1) use closed population models (e.g., Lincoln-Peterson) to estimate population size for short term studies where closure assumption can be met, (2) use the robust desi...

Modeling capture, recapture, and removal statistics for estimation of demographic parameters for fish and wildlife populations : past, present, and future

Abstract In this article I review the modeling of capture, recapture, and removal statistics for the purpose of estimating demographic parameters of fish and wildlife populations. Topics considered include capture-recapture models, band or tag return models, removal and catch per unit effort models, selective removal or change-in-ratio models, radio-tagging survival models, and nest survival models. The purpose is to present important concepts in a general manner for the benefit of a wide audience of statisticians. I will not attempt to be comprehensive, and I indulge in speculation about future directions. I indicate the importance of different statistical tools to this subject, such as Bayesian inference, “boot strapping,” robustness studies, goodness-of-fit tests. I also emphasize connections to other application areas of statistics. Capture-recapture methods, for example, are being considered for estimation of a variety of elusive human populations, such as the homeless and people missed in the census...

Estimating Breeding Proportions and Testing Hypotheses about Costs of Reproduction with Capture-Recapture Data

The proportion of animals in a population that breeds is an important determinant of population growth rate. Usual estimates of this quantity from field sampling data assume that the probability of appearing in the capture or count statistic is the same for animals that do and do not breed. A similar assumption is required by most existing methods used to test ecologically interesting hypotheses about reproductive costs using field sampling data. However, in many field sampling situations breeding and nonbreeding animals are likely to exhibit different probabilities of being seen or caught. In this paper, we propose the use of multistate capture-recapture models for these estimation and testing problems. This methodology permits a formal test of the hypothesis of equal capture/ sighting probabilities for breeding and nonbreeding individuals. Two estimators of breeding proportion (and associated standard errors) are presented, one for the case of equal capture probabilities and one for the case of unequal capture probabilities. The multistate modeling framework also yields formal tests of hypotheses about reproductive costs to future repro- duction or survival or both fitness components. The general methodology is illustrated using capture-recapture data on female meadow voles, Microtus pennsylvanicus. Resulting estimates of the proportion of reproductively active females showed strong seasonal vari- ation, as expected, with low breeding proportions in midwinter. We found no evidence of reproductive costs extracted in subsequent survival or reproduction. We believe that this methodological framework has wide application to problems in animal ecology concerning breeding proportions and phenotypic reproductive costs.

Estimation methodology in contemporary small mammal capture-recapture studies

Estimators of population size and survival rate based on the Jolly-Seber capture-recapture model and the “enumeration method” are described. Enumeration estimators are shown to estimate complicated functions of capture and survival probabilities and, in the case of the population size estimator, population size. Frequently-listed reasons for preferring enumeration estimators are discussed and the Jolly-Seber estimators are shown to be superior even in the case of heterogeneity and trap-happy response, the two sources of unequal capture probability most likely to occur in small mammal studies. New developments in probabilistic capture-recapture models are described, and these models are recommended for future small mammal capture-recapture studies.

Forest fragmentation and bird community dynamics: inference at regional scales

With increasing fragmentation of natural areas and a dramatic reduction of forest cover in several parts of the world, quantifying the impact of such changes on species richness and community dynamics has been a subject of much concern. Here, we tested whether in more fragmented landscapes there was a lower number of area-sensitive species and higher local extinction and turnover rates, which could explain higher temporal vari- ability in species richness. To investigate such potential landscape effects at a regional scale, we merged two independent, large-scale monitoring efforts: the North American Breeding Bird Survey (BBS) and the Land Use and Land Cover Classification data from the U.S. Geological Survey. We used methods that accounted for heterogeneity in the probability of detecting species to estimate species richness and temporal changes in the bird communities for BBS routes in three mid-Atlantic U.S. states. Forest breeding bird species were grouped prior to the analyses into area-sensitive and non-area-sensitive species according to previous studies. We tested predictions relating measures of forest structure at one point in time (1974) to species richness at that time and to parameters of forest bird community change over the following 22-yr-period (1975-1996). We used the mean size of forest patches to characterize landscape structure, as high correlations among landscape variables did not allow us to disentangle the relative roles of habitat fragmentation per se and habitat loss. As predicted, together with lower species richness for area-sensitive species on routes surrounded by landscapes with lower mean forest-patch size, we found higher mean year- to-year rates of local extinction. Moreover, the mean year-to-year rates of local turnover (proportion of locally new species) for area-sensitive species were also higher in landscapes with lower mean forest-patch size. These associations were not observed for the non-area- sensitive species group. These results suggest that landscape structure may influence forest bird communities at regional scales through its effects on the total number of species but also on the temporal rates of change in community composition. Evidence for higher rates of local extinction and turnover in more fragmented landscapes suggests that bird communities function as metapopulations at a regional scale, and points out the importance of colonizations and recolonizations from surrounding landscapes to local community dynamics. Further, our results illustrate that the methods used to estimate the community parameters can be a powerful statistical tool in addressing questions relative to the dynamics of communities.